Article

Maximum polyphyly: Multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae)

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Abstract

Minuartia is one of the larger genera of Caryophyllaceae with about 175 species distributed mainly in the Northern Hemisphere. The taxonomy of the genus has been studied by several authors, resulting in the recognition of numerous infrageneric groups. Previous phylogenies of the Caryophyllaceae showed that Minuartia is polyphyletic, but included only a limited sample of the genus. We here provide a molecular phylogeny of Minuartia including all infrageneric groups recognized in the last revision of the genus. We reconstructed the phylogeny of the genus using DNA sequences of nrITS and plastid matK from 160 ingroup and 105 outgroup samples. The evolution of the morphological character relevant for the delimitation of Minuartia (three styles plus three capsule valves/teeth) was reconstructed. Minuartiahas been defined with a plesiomorphic character and is highly polyphyletic. All four subgenera fall into different lineages containing other genera of the family, and M. subg. Minuartia, as by far the largest subgenus, falls into seven clades, which together do not form a monophylum. These clades are closely related to several other genera, e.g., Sagina, Colobanthus and Scleranthus. In several cases taxonomic groups below subgeneric rank are monophyletic. The type of Minuartia, M. dichotoma, is part of a clade containing M. sect. Plurinerviae and sect. Minuartia. We propose to retain this clade as Minuartia s.str. and to transfer the other species of Minuartia to the genera Cherleria, Eremogone, Facchinia, Mcneillia, Minuartiella, Mononeuria, Pseudocherleria, Rhodalsine, Sabulina and Triplateia.

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... Consequently, a new subfamily named Eremogoneae Rabeler & W.L.Wagner was added to the subfamilies Alsinoideae Burnett, Caryophylloideae Arn. and Paronychioideae A.St. (Bittrich 1993;Harbaugh et al. 2010). In addition, several studies have been conducted to determine the phylogeny of the genera Arenaria L., Minuartia L., Dianthus L. (ca 300 sp.), Gypsophila L. (ca 150 sp.) and Silene L. As a result, many new genera have been described, while others have become synonymous (Dillenberger and Kadereit 2014;Sadeghian et al. 2015;Madhani et al. 2018). ...
... The genus Minuartia differs morphologically from closely related genera in having three styles and capsule teeth as many as styles (McNeill 1963(McNeill , 1967. However, molecular studies have shown that the genus is polyphyletic (Minuto et al. 2011;Harbaugh et al. 2010;Greenberg and Donoghue 2011;Dillenberger and Kadereit 2014). For this reason, Dillenberger and Kadereit (2014) analysed cpDNA and nrDNA sequence data and found the sampled taxa forming an isolated clade that could be interpreted as sister to a clade that included Colobanthus Bartl., Facchinia Rchb., Sabulina Rchb. ...
... However, molecular studies have shown that the genus is polyphyletic (Minuto et al. 2011;Harbaugh et al. 2010;Greenberg and Donoghue 2011;Dillenberger and Kadereit 2014). For this reason, Dillenberger and Kadereit (2014) analysed cpDNA and nrDNA sequence data and found the sampled taxa forming an isolated clade that could be interpreted as sister to a clade that included Colobanthus Bartl., Facchinia Rchb., Sabulina Rchb. and Sagina L. Consequently, they accepted eleven minor genera segrated from Minuartias.l., on top of them Sabulina (Rchb.) ...
Article
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The genus Sabulina is represented by 26 taxa in Turkey. Some interesting specimens belonging to the genus, the main distribution area of which is the Mediterranean phytogeographical region, were collected in Erzurum province. These newly collected specimens and Sabulina rimarum var. rimarum were compared with each other in terms of their general morphology and seed micromorphology. Besides, by using DNA sequences of ITS genes, phylogenetic relationships between these collected species and the other species were investigated. As a result of the evaluation of molecular and morphological data together, it was decided that the specimens collected from Erzurum belong to a new species for the science world. This species was named Sabulina tortumensis, and its description, pictures, distribution, habitat preferences, degree of endangerment and phylogenetic relationships with closely related species are presented. Additionally, we propose new combinations for five taxa of Sabulina that did not have Sabulina names and provide a key to annual species of the genus in Turkey.
... In Sagina, outer stamens and capsule valves are episepalous and inner stamens and styles are alternisepalous (Crow 1978;Timaná 2018). Although the most comprehensive phylogeny including Colobanthus and Sagina revealed the two genera to be monophyletic, as assumed by Bittrich (1993), it contained only four species of each genus (Dillenberger and Kadereit 2014). Although Pufal (2010) analyzed a larger species sample of Colobanthus, the sequences used are not publicly available. ...
... The closest relatives of Sagina and Colobanthus, i.e. Facchinia and Sabulina Rchb., (Dillenberger and Kadereit 2014) are predominantly Northern Hemisphere genera with together more than 80 species. Two other genera, Bufonia L., and Drypis L., possibly also belonging to tribe Sagineae sensu Harbaugh et al. (2010), are found only in the Northern Hemisphere. ...
... These DNA regions had proved to be phylogenetically informative in Caryophyllaceae in previous studies (e.g. Dillenberger and Kadereit 2014;Zhang et al. 2016;Legler and Dillenberger 2017;Biersma et al. 2020). Polymerase chain reactions (PCR) for ITS and trnQ-rps16 were carried out as described in Kadereit (2014, 2017) and Biersma et al. (2020). ...
Article
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Colobanthus (23 species) and Sagina (30–33 species) together are sister to Facchinia. Whereas Facchinia is distributed in western Eurasia, Colobanthus is almost exclusively distributed in the Southern Hemisphere, and Sagina is distributed in both hemispheres with the highest species diversity in western Eurasia. We examined: 1. Whether Sagina and Colobanthus are monophyletic sister genera, 2. Where the two genera originated and how many times dispersal between hemispheres occurred, and 3. Which colonization routes between hemispheres were taken. We reconstructed the phylogeny of Colobanthus and Sagina using nuclear ribosomal internal transcribed spacer (ITS) and two plastid spacers (cpDNA) of altogether 158 ingroup samples of 45 species, and performed molecular dating and ancestral area reconstructions. Sagina and Colobanthus were confirmed as monophyletic sister genera. Biogeographical reconstructions based on ITS and cpDNA showed that Sagina reached the Southern Hemisphere in Australasia or in Africa. For Colobanthus, patterns were less clear and less well-supported: ITS showed Australasia as the region of entry, but cpDNA implied that the Southern Hemisphere may have been entered in America. The extant distributions and the biogeographical histories of Colobanthus and Sagina show both similarities and dissimilarities. This illustrates that biogeographical histories, even of closely related and ecologically very similar lineages, can be highly idiosyncratic.
... . Minuartia has been proven to not be a monophyletic group, and its infrageneric classification is inconclusive (L€ ove and L€ ove 1975;Dillenberger and Kadereit 2014). ...
... Recent studies of phylogenetic relationships indicate that Arenaria is not a monophyletic group and subgenus Odontostemma could be clustered into a monophyletic group, which would support restoring the subgenus Odontostemma to Odontostemma Harbaugh et al. 2010;Greenberg and Donoghue 2011;Dillenberger and Kadereit 2014;Sadeghian et al. 2015). This view has gained the support of many scholars (Hern andez-Ledesma et al. 2015; Richard and Wagner 2016; Yao 2017). ...
... However, phylogenetic relationships revealed in recent years indicate that Minuartia is not a monophyletic group Harbaugh et al. 2010;Greenberg and Donoghue 2011), and some sections in the subgenus Minuartia have also been proven not to be monophyletic. For example, section Spectabiles is embedded in different clades in the phylogenetic tree (Dillenberger and Kadereit 2014). ...
Article
Pollen morphology of 79 species, one subspecies and eight varieties representing nine tribe Alsineae and two tribe Sperguleae genera was studied using light microscopy (LM) and scanning electron microscopy (SEM). Among them, 53 species and eight varieties (22 species of which are endemic to China) were reported for the first time. The results demonstrate that the number of pores, pollen surface ornamentation and microechini density have important systematic significance. The surface ornamentation can be divided into three types, namely microechinate-perforate, microechinate-punctate, and microechinate-punctate-perforate. Pollen characteristics support: (i) isolation of Arenaria subgenus Odontostemma from Arenaria and reclassification of Arenaria subgenus Odontostemma to Odontostemma; (ii) isolation of Cerastium subgenus Dichodon from Cerastium and reclassification of Cerastium subgenus Dichodon to Dichodon; (iii) isolation of Minuartia subgenus Rhodalsine from Minuartia and reclassification of Minuartia subgenus Rhodalsine to Rhodalsine; (iv) isolation of Rhodalsine from tribe Alsineae and reclassification of Rhodalsine into tribe Sperguleae; and (v) isolation of Sagina from tribe Alsineae. In addition, the relationships of Arenaria subgenus Eremogone and Arenaria subgenus Eremogoneastrum to tribe Alsineae need to be further studied. Furthermore, pollen characteristics indicate that tribe Alsineae is more evolved than tribe Sperguleae.
... 150 sp.), Minuartia Linnaeus (1753: 89), Polycarpon Linnaeus (1759: 859, 881), Silene Linnaeus (1753: 416), ect. (see e.g., Kool et al. 2007, Dillenberger & Kadereit 2014, Iamonico 2013, 2015, Iamonico & Domina 2015, Madhani et al. 2018, Sadeghian et al. 2015. ...
... From the morphological point of view, the genus Minuartia differs from the related ones by having three styles and capsule teeth as many as styles (see e.g., McNeill 1963, 1967, Dillenberger & Kadareit 2015, Koç et al. 2019. Phylogenetic studies have shown that the genus is highly polyphyletic , Greenberg et al. 2011, Dillenberger & Kadereit 2014. Dillenberger & Kadereit (2014) made a phylogeny of Minuartia using cpDNa and nrDNa sequence data and proposed 10 segregated genera which were subsequently accepted by several authors (e.g., Conti et al. 2014, iamonico 2014, Legler & Dillenberger 2017, Moore & Dillenberger 2017, Dillenberger & rabeler 2018. ...
... Phylogenetic studies have shown that the genus is highly polyphyletic , Greenberg et al. 2011, Dillenberger & Kadereit 2014. Dillenberger & Kadereit (2014) made a phylogeny of Minuartia using cpDNa and nrDNa sequence data and proposed 10 segregated genera which were subsequently accepted by several authors (e.g., Conti et al. 2014, iamonico 2014, Legler & Dillenberger 2017, Moore & Dillenberger 2017, Dillenberger & rabeler 2018. among these genera there is Minuartiella Dillenberger & Kadereit (2014: 78) which currentl includes four species [previously belonging to Minuartia sect. ...
Article
The genus Minuartiella comprises four species distributed in the E-Mediterranean region and SW-Asia. Some interesting specimens belonging to the genus where collected from Burdur province (Turkey). On the basis of macro- and micromprhological analyses, as well as molecular data, a new species—Minuartiella serpentinicola sp. nov.—and a new combination—Minuartiella ×antalyensis comb. nov.—are proposed in the present paper. A detailed description of the new species as well as pictures, distribution map, habitat, and IUCN category are also provided. A diagnostic key of Mnuartiella species is provided.
... s.l. [26]. ...
... Dillenberger and Kadereit [26] documented rampant polyphyly in Minuartia as traditionally conceived, revealing that the main diagnostic characters for the genus (i.e., presence of three styles and three fruit valves) are indeed plesiomorphic. Even subgeneric ranks (e.g., Minuartia subg. ...
... A synopsis of the most relevant taxonomic treatments is provided in Table 1. In addition, the relationships between the taxa are still largely speculative (e.g., [27]), and the phylogeny almost unknown; as at present only four species (mostly with single specimens) have been investigated by molecular methods [26]. In this study, we infer nuclear and plastid phylogenies of genus Mcneillia involving all its species and subspecies across their geographic distributions. ...
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The genus Mcneillia has been recently segregated from Minuartia L. based on molecular results, also supported by morphology. However, to date, a comprehensive study on the phylogenetic relationships within this genus is lacking. In this paper, we provide a multigene phylogeny of all the species and subspecies of Mcneillia employing two nuclear and six chloroplast markers. We documented extensive gene flow between taxa, sometimes separated at specific rank. In addition, Mcneillia as currently circumscribed, is not monophyletic. In fact, Mcneillia graminifolia subsp. brachypetala, strictly endemic to Greece, truly belongs to Minuartiella, a genus otherwise limited to South-West Asia. Moreover, even after removal of this taxon, our results do not support the monophyly of the taxa included in M. graminifolia s.l., the most variable and widespread species of the genus. Further controversial subspecies of Mcneillia graminifolia, i.e., subsp. hungarica and subsp. rosanoi, are shown to deserve taxonomic recognition as separate species, whereas Mc. moraldoi is not distinct at specific rank. In addition, Mc. saxifraga subsp. tmolea is here regarded as a further distinct species. A consistent taxonomic treatment is therefore proposed with six new combinations and nomenclatural notes, providing the necessary typifications.
... collina (Neilr.) Domin) -stony slopes -C -The nomenclature according to Dillenberger and Kadereit (2014). Sabulina mediterranea (Ledeb. ...
... (Conti 1995). The nomenclature according to Dillenberger and Kadereit (2014). Sabulina tenuifolia (L.) Rchb. ...
... subsp. hybrida) -arid meadows -C -The nomenclature according to Dillenberger and Kadereit (2014). Juniperus sabina L. -montane stony slopes -R -Barrea at Colle S. Angelo, near Villavallelonga!, Monte Boccanera, Mainarde on Morrone delle Rose! D'Andrea 1982;Conti et al. 1987;Rovelli 1992). ...
... while Gucel (2013) recorded 2n=30 in M. nifensis. Dillenberger & Kadereit (2013) reported 2n=30 in M. hamata and 2n= 22, 46 and 48 in M. hybrida. Dillenberger & Kadereit (2013) reported values for M. meyeri (2n=30), M. montana (2n=56), M. recurva (2n=30) and M. picta ( 2n=22). ...
... Dillenberger & Kadereit (2013) reported 2n=30 in M. hamata and 2n= 22, 46 and 48 in M. hybrida. Dillenberger & Kadereit (2013) reported values for M. meyeri (2n=30), M. montana (2n=56), M. recurva (2n=30) and M. picta ( 2n=22). Ghaffari & Kelich (2006) studied M. lineate and reported a value of n=10. ...
... Our cytological studies are based on six taxa comprising Minuartia. M. picta are showing the dibasic nature of the genus with x=11 and 14, this result agreed with the findings of Dillenberger & Kadereit (2013) who recorded a value of x=11, and our In this study, the chromosome numbers, karyotypes, ideograms, and karyotype asymmetry degrees of Minuartia were determined. ...
Article
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Chromosome count, karyotypic character analysis, meiotic studies, monoploid karyograms and ideograms were performed in six taxa of Minuartia growing in Iraq (M. hamata, M. hybrida subsp. hybrida, M. intermedia, M. meyeri, M. picta and M. hybrida subsp. turcica). Species of M. hamata and M. meyeri showed 2n=2x=30 chromosome number, while M. hybrida subsp. hybrida and M. intermedia were diploid (26). The chromosome number (n=x) of six species was studied, and was found to be n=15 in M. hamata and M. meyeri, 13 in M. hybrida and M. intermedia, while in M. picta we recorded values of n= 11 and 14. Karyotype analysis of this species was first carried out in our study. Analysis of metaphases showed that the karyotype formula was mainly metacentric, submetacentric, and sub acrocentric. The sizes of the chromosomes were mainly small and very small. The course of meiosis varied from normal to abnormal. Abnormal microsporogenesis formation of two bridge chromosomes was detected in M. hamata and one bridge chromosome in M. intermedia and M. meyeri. Formation of laggard's chromosomes was detected in M. hamata, M. meyeri and M. intermedia. As well as ring chromosome was showed in M. hybrida subsp. hybrida, also, some cells contain triad cell in metaphase stage instead four cells, as well as founded cell, contains two nuclei in same species which led to reduced pollen fertility and differences in pollen grain size.
... 150 sp.), Polycarpon Linnaeus (1759: 881) s.l., Silene Linnaeus (1753: 416), ect. (see e.g., Dillenberger & Kadereit 2014, Iamonico 2013, 2015, Iamonico & Domina 2015, Madhani et al. 2018, Sadeghian et al. 2015. ...
... accoding to McNeill (1963McNeill ( , 1967 the genus Minuartia would morphologically differs from close related genera by having three styles and capsule teeth as many as styles. however, molecular studies have shown that the genus is highly polyphyletic, and 11 different genera were considered by Dillenberger & Kadereit (2014), Greenberg &Donoghue (2011), andharbaugh et al. (2010), and subsequent accepted by several authors (e.g., Conti et al. 2014, iamonico 2014, Legler & Dillenberger 2017, Moore & Dillenberger 2017, Dıllenberger & rabeler 2018. ...
... 15 Minuartia specimens from different populations growing in antalya province were collected. among these specimens, those growing in the a stony area with serpentines attracted were compared with the related ones, based on literature (Bojnansky & Fargasová 2007, Dillenberger & Kadereit 2014, Mostafavi et al. 2013, Parolly & eren 2007, tan & Vural 2000, McNeill 1963, 1967, Shishkin 1995, rechinger 1964, halliday 1976, Zohary 1966, Kamari 1997) and the examination of specimens kept in herbaria GaZI, aNK, and huB; specimens found in K, G, Je and e virtual herbaria were also examined (acronyms according to thiers 2019-onward). ...
Article
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The genus Minuartia is represented in Turkey by 34 taxa. Some interesting specimens were collected from Antalya province, and examined. These specimens resemble Minuartia meyeri, and M. multinervis from which differ by characters (macro-, and micromorphological) of inflorescence, alar pedicels, petals, sepals, capsules and seeds. Moreover, by using the DNA sequences of the ITS genes, phylogenetic relationships between this collected species, and the related species were investigated. As a result of the evaluation of molecular, and morphological data, we proposed to described the population from Antalya as a new species for the science. A description, pictures, distribution, habitat, and IUCN category are given.
... Arenaria (Sadeghian & al., 2015) and Minuartia Loefl. (Dillenberger & Kadereit, 2014). Sadeghian & al. (2015) confirmed the exclusion of the majority of Arenaria spp. ...
... Odontostemma to genus rank, maintaining them as members of Alsineae. Minuartia, also treated as a member of Alsineae in traditional systems (Pax & Hoffmann, 1934;Schischkin, 1936;Bittrich, 1993), turned out to be polyphyletic with members placed in both tribes Sagineae J.Presl and Sclerantheae DC. (Dillenberger & Kadereit, 2014). A recent study of Pseudostellaria retrieved this genus as a member of the Odontostemma clade, and the new genus Hartmaniella M.L.Zhang & Rabeler was proposed for two American species formerly assigned to Pseudostellaria (Zhang & al., 2017). ...
... -The placement of a few species of Stellaria, i.e., St. ovata (Mexico to S America), St. minutifolia, and St. howardii (Dominican Republic), among the outgroups is in agreement with previous molecular phylogenies (Harbaugh & al., 2010;Greenberg & Donoghue, 2011;Dillenberger & Kadereit, 2014). Morphological and geographic evidence (growth habit, 4-merous flowers, relatively shallowly cleft petals, seed with prominent dorsal tubercles, and distribution in Antillean islands) suggests St. antillana Urb. is closely allied to St. ovata (Maguire, 1958) and belongs to this group, which was also confirmed by a recent phylogenetic analysis (Sharples & Tripp, 2019a). ...
Article
The tribe Alsineae is a large monophyletic group in the family Caryophyllaceae especially found across Eurasia and the Americas, but with a center of diversity in the Mediterranean region. Several previous molecular phylogenetic studies have focused on the delimitation of genera and tribes of Caryophyllaceae, especially the subfamily Alsinoideae or the tribe Alsineae in a broader sense than now recognized. However, there are still many open questions regarding the subdivision of the tribe and genus delimitation. In the present study, we sampled 191 (148 species) and 149 (125 species) accessions of Alsineae representing almost all (Adenonema and Pseudocerastium were not available to us) recognized genera in the tribe for nuclear DNA internal transcribed spacer (ITS) and plastid marker rps16 sequences, respectively. A combined matrix of 103 species was built for taxa with both sequences available. Maximum parsimony and Bayesian inference analyses retrieved Cerastium and Stellaria (including Myosoton) as the largest monophyletic genera, while other genera were medium‐sized (10–20 spp.) or small (<10 spp.). Our expanded sampling of Pseudostellaria and its relatives suggests a broader circumscription of this genus. Major divergence in morphology, particularly of the seeds, observed in the “Protostellaria”‐clade, allows recognition of some taxonomic changes. A total of 16 genera are recognized in Alsineae including Cerastium, Dichodon, Hartmaniella, Holosteum, Lepyrodiclis, Mesostemma, Moenchia, Nubelaria, Odontostemma, Pseudostellaria, Rabelera, Schizotechium, Shivparvatia, and Stellaria, along with Adenonema and Pseudocerastium that could not be analyzed and are, therefore, kept as distinct genera. A diagnostic key to these genera, as well as notes on their relationships, distribution, and nomenclature, is provided.
... The genus Minuartia L. (Family: Caryophyllaceae) is the particularly of interest because of its preference for metalliferous and other extreme substrates. This genus, previously included in Arenaria (Maguire 1951), comprises > 170 species distributed mainly in the Northern Hemisphere, between temperate and arctic-alpine region (Dillenberger and Kadereit 2014). One of the main centers of Minuartia diversity is Europe (85 taxa), especially the Mediterranean region (Halliday 1993;Diklić and Stevanović 2012), with a significant number of species in the Balkan Peninsula (47 taxa; Diklić and Stevanović 2012). ...
... Graebn., generally calcifuge species from the monophyletic Sect. Plurinervie McNeill (Dillenberger and Kadereit 2014). Although also considered a subspecies (M. ...
... Stoj. & Stef.; Dillenberger and Kadereit 2014), M. bulgarica is an accepted taxon according to relevant databases (Euro+Med 2006; http:// www. thepl antli st. ...
Article
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The aim of this study was to determine the concentrations of potentially toxic elements in soil samples and plant tissues of Minuartia recurva and M. bulgarica, predominantly or exclusively calcifuge species. Biological concentration (BCs) and translocation factors (TFs) were used to evaluate their accumulation potential. Considerable differences were observed between M. recurva and M. bulgarica assessions in terms of accumulation strategies of potentially toxic elements (PTEs). In M. recurva, most of the elements analyzed (Mn, Cu, Zn, Cd, and Co) were transported to the shoot, whereas in M. bulgarica, these elements remained predominantly in the roots. The Cu concentrations in the shoot samples of M. recurva from an abandoned iron-copper mine at Mt. Kopaonik were clearly above the notional hyperaccumulation threshold, characterizing this species as a possible Cu hyperaccumulator. Additionally, strong accumulation potential for Cr, Ni, Zn, Pb, and Cd was observed in M. recurva assessions, but without significant accumulation due to the low concentrations of these elements in the soils. The strong accumulation capacity and the different strategies in tolerance to PTEs indicate a potential of the two species for an application in phytoremediation: M. recurva for phytoextraction and M. bulgarica for phytostabilization.
... 30 annual or perennial herbaceous to small subshrubby species growing mostly on dry gravely slopes in mountainous regions (bittrich 1993, Chrtek & Křísa 1999 and is distributed in the Mediterranean and the Irano-turanian regions (see bittrich 1993). the genera of Caryophyllaceae are under partial rearrangements and several genera need further study from both taxonomical and nomenclatural points of view (see e.g., Dillenberger & Kadereit 2014, Iamonico 2015, 2016a, 2016b, 2018, Iamonico & Domina 2015. a subfamilial classification of Caryophyllaceae is rejected by the molecular phylogenetic investigations and a tribal system is currently followed (harbaugh et al. 2010, Greenberg & Donoghue 2011, hernández-Ledesma et al. 2015. ...
... Cerastium Linnaeus (1753: 437) and Sagina Linnaeus (1753: 128) have traditionally been considered as close relatives of Bufonia (Chrtek & Křísa 1999). however, molecular phylogenetic studies suggested Sabulina reichenbach (1832: 24) and Sagina as the closest allies of Bufonia (Dillenberger & Kadereit 2014, Greenberg & Donoghue 2011). In addition, major morphological similarities between Bufonia and Eremogone fenzl (1833: 13) species were reported by Sadegian et al. (2015). ...
... In the present study, we perform a comparative study of stem anatomy in Bufonia and its nine related genera based on the recent molecular phylogenetic findings (harbaugh et al. 2010;Greenberg & Donoghue 2011, Dillenberger & Kadereit 2014, hernandez-Ledesma et al. 2015, Sadeghian et al. 2015), i.e., Arenaria, Cerastium, Eremogone, Minuarta, Minuartiella Dillenberger & Kadereit (2014, Odontostemma bentham ex Don (1831: 449), Shivparvatia Pusalkar & D.K.Singh (2015: 81), Sabulina, and Sagina. the aims of this study are: 1) to evaluate the usefulness of stem anatomy in generic delimitation of Bufonia, 2) to elucidate potential implication of anatomical characters in infrageneric classification of Bufonia, and 3) to find evidence supporting the suggestecd phylogenetic placement of taxa related to or assigned once to Bufonia. ...
Article
Stem anatomy of Bufonia and nine allied genera was studied in order to investigate the systematic value of anatomical characters in generic delimitation. Cross-sections of stems of 31 accessions representing 14 species and 3 subspecies of Bufonia, and 17 additional species of the genera Arenaria, Cerastium, Eremogone, Minuartia, Minuartiella, Odontostemma, Sabulina, Sagina, and Shivparvatia were examined using light microscopy. The characteristic features of stem anatomy in Bufonia are: 1) presence of one to four layers of parenchyma in cortex, 2) calcium oxalate crystals mostly in the endodermis, 3) presence of inner and outer sclerenchymatous pericycle, and 4) a large central pith rupturing in a linear to boat-shape mode and forming a central continuous slit. Characters such as arrangement and continuity of xylem along with the number, and arrangement of different tissues (or layers) discriminate Bufonia from the other studied genera. Stem anatomy of Eremogone, Minuartia, and Bufonia showed some degree of similarity reflecting a level of convergence in the stem anatomy that might explain the taxonomic confusion associated with this group. The present study supports previous exclusion of Bufonia koelzii and B. caespitosa from Bufonia and their inclusion in Eremogone.
... while Gucel (2013) recorded 2n=30 in M. nifensis. Dillenberger & Kadereit (2013) reported 2n=30 in M. hamata and 2n= 22, 46 and 48 in M. hybrida. Dillenberger & Kadereit (2013) reported values for M. meyeri (2n=30), M. montana (2n=56), M. recurva (2n=30) and M. picta ( 2n=22). ...
... Dillenberger & Kadereit (2013) reported 2n=30 in M. hamata and 2n= 22, 46 and 48 in M. hybrida. Dillenberger & Kadereit (2013) reported values for M. meyeri (2n=30), M. montana (2n=56), M. recurva (2n=30) and M. picta ( 2n=22). Ghaffari & Kelich (2006) studied M. lineate and reported a value of n=10. ...
... Our cytological studies are based on six taxa comprising Minuartia. M. picta are showing the dibasic nature of the genus with x=11 and 14, this result agreed with the findings of Dillenberger & Kadereit (2013) who recorded a value of x=11, and our In this study, the chromosome numbers, karyotypes, ideograms, and karyotype asymmetry degrees of Minuartia were determined. ...
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Chromosome count, karyotypic character analysis, meiotic studies, monoploid karyograms and ideograms were performed in six taxa of Minuartia growing in Iraq (M. hamata, M. hybrida subsp. hybrida, M. intermedia, M. meyeri, M. picta and M. hybrida subsp. turcica). Species of M. hamata and M. meyeri showed 2n=2x=30 chromosome number, while M. hybrida subsp. hybrida and M. intermedia were diploid (26). The chromosome number (n=x) of six species was studied, and was found to be n=15 in M. hamata and M. meyeri, 13 in M. hybrida and M. intermedia, while in M. picta we recorded values of n= 11 and 14. Karyotype analysis of this species was first carried out in our study. Analysis of metaphases showed that the karyotype formula was mainly metacentric, submetacentric, and sub acrocentric. The sizes of the chromosomes were mainly small and very small. The course of meiosis varied from normal to abnormal. Abnormal microsporogenesis formation of two bridge chromosomes was detected in M. hamata and one bridge chromosome in M. intermedia and M. meyeri. Formation of laggard’s chromosomes was detected in M. hamata, M. meyeri and M. intermedia. As well as ring chromosome was showed in M. hybrida subsp. hybrida, also, some cells contain triad cell in metaphase stage instead four cells, as well as founded cell, contains two nuclei in same species which led to reduced pollen fertility and differences in pollen grain size.
... Greenberg and Donoghue 2011). At genus rank, several studies have been carried out on Arenaria L., Minuartia L., Dianthus L., Gypsophila L., Polycarpon L., Silene L., etc. (see e.g., Kool et al. 2007, Iamonico 2013, 2015, 2016, Dillenberger and Kadereit 2014, Iamonico and Domina 2015, Sadeghian et al. 2015, Dillenberger and Rabeler 2018, Madhani et al. 2018), but various questions are still open. ...
... This clade corresponds to the tribe Sagineae. Dillenberger and Kadereit (2014), who investigated in detail the genus Minuartia, did not consider the genus Habrosia in their analysis. However, the species of Minuartia, included in the tribe Sagineae by Greenberg and Donoghue (2011), were treated by Dillenberger and Kadereit (2014) as belonging to the resurrected genus Sabulina Rchb. ...
... Dillenberger and Kadereit (2014), who investigated in detail the genus Minuartia, did not consider the genus Habrosia in their analysis. However, the species of Minuartia, included in the tribe Sagineae by Greenberg and Donoghue (2011), were treated by Dillenberger and Kadereit (2014) as belonging to the resurrected genus Sabulina Rchb. In contrast to Habrosia, Dillenberger and Kadereit (2014) included Arenaria fontinalis in their analysis, which was also investigated by Greenberg and Donoghue (2011), and confirmed that it is to be treated as a member of Sabulina and, in fact, a new combination, S. fontinalis (Short and R. Peter) Dillenb. ...
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Habrosia (Sagineae, Caryophyllaceae) is a genus that includes only H. spinuliflora, a species occurring in Iran, Iraq, Syria, Lebanon, and Turkey (Irano-Turanian floristic chorological element). Based on the available molecular data published in 2011, Habrosia appears to be nested in a Minuartia-clade, which includes taxa currently recognized under the genus Sabulina. Consequently, Habrosia should be treated as a genus to be included in Sabulina. However, the molecular tree published in 2011 considered only 9 Sabulina members whereas, according to the current concept, Sabulina is a genus comprising about 65 species. Unfortunately, the molecular phylogeny including a larger Sabulina sample published in 2014 did not include H. spinuliflora and the taxonomic position of Habrosia remains, therefore, uncertain. With the aim of verifying the correct position of Habrosia in the tribe Sagineae with respect to its relationship to Sabulina, a comprehensive molecular investigation based on ITS sequences, linked to detailed morphological data, is presented. The results obtained revealed that Habrosia is not part of Sabulina. A detailed description of H. spinuliflora, its ecological preference, and a distribution map are provided. Eventually, the name Arenaria spinulifolia (basionym of H. spinuliflora) is lectotypified on a specimen preserved at G (barcode G00212963).
... Ancestral range reconstruction. -Based on present comprehensive systematic literature pertaining to Caryophyllaceae (Greenberg & Donoghue, 2011;Dillenberger & Kadereit, 2014;Sadeghian & al., 2015), the distributions of nearly 500 taxa with available DNA sequences were checked through the Global Biodiversity Information Facility (GBIF, https://www.gbif.org/), and we delineated nine geographical areas for ancestral range reconstruction: ( Ancestral range reconstruction and estimation of spatial patterns of geographic diversification were undertaken using the Bayesian Binary MCMC (BBM) method implemented in RASP (Reconstruct Ancestral State in Phylogenies) v.3.2 (Yu & al., 2015) and a maximum likelihood-based DEC (dispersal-extinction-cladogenesis) model in Lagrange v.20130526 (Ree & Smith, 2008). ...
... Arenaria taibaishanensis shares some morphological characteristics with M. biflora, including infertile axial foliage clusters, apiculate and narrowly ovoid/ovoid-cylindric capsules which are 3-valved and protrude from the sepals during frutescence (Fig. 1L). Minuartia, a highly polyphyletic genus, has been broken up into 11 different genera based on recent molecular phylogenetic studies (Harbaugh & al., 2010;Greenberg & Donoghue 2011;Dillenberger & Kadereit 2014). Minuartia biflora was transferred to Cherleria L. (about 20 species) and named C. biflora (L.) A.J.Moore & Dillenb. ...
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Cushion plants exhibiting adaptive convergence in cold and dry environments are keystone and foundation species in the alpine/subnival habitat on the Qinghai‐Tibet Plateau (QTP). To date, little attention has been paid to the molecular phylogeny, origin, and biogeography of cushion plants on the QTP. We investigated the molecular phylogeny of classic cushion Arenaria subg. Dolophragma, A. subg. Eremogoneastrum, and Thylacospermum on the QTP, within the framework of the Caryophyllaceae. A new Thylacospermum‐Dolophragma clade was identified using combined plastid markers (rps16, matK, trnL‐trnF, trnS‐trnfM) and nuclear ribosomal DNA. Molecular divergence dating suggested that Thylacospermum and A. subg. Dolophragma originated in the middle to late Miocene (11.68 Ma). Combined with ancestral range reconstruction, there is an indication that all species of cushion Arenaria (A. subg. Dolophragma, A. subg. Eremogoneastrum) originated during/after the late Pliocene, and the QTP was their ancestral area. Furthermore, ecological niche modeling showed that the areas occupied by the three studied cushion plants during the last glacial maximum (LGM) was broader than that of their present distribution, implying a reduction in their range after the LGM. This evidence clearly illustrates that multistage uplift of the QTP and bordering mountains since the Miocene associated with climatic change (worldwide cooling, aridification in Central Asia, Quaternary glaciation) played a role in triggering and facilitating the speciation and/or evolutionary radiations of the species studied.
... The taxonomy of the group is in a state of flux with controversies surrounding the delimitation of taxa and the designation of their ranks (Graebner 1918;Hayek 1922;Pawłowski 1939;Halliday 1964Halliday , 1993Kamari and Constantidinis 1994;Dvořáková 2000). Recently, Dillenberger and Kadereit (2014) revealed the genus Minuartia to be highly polyphyletic and re-established the separate genus Sabulina, comprising ca 65 species in the proposed circumscription, including the Sabulina (Minuartia) verna group. The phylogeny of the group (10 accessions assigned to 6 species) nevertheless remains unresolved. ...
... Our results also at least partly support recent concepts adopted in Dillenberger and Kadereit (2014) ...
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Polyploidization, a key driver of plant diversification, is believed to have interacted with Pleistocene climatic oscillations and local ecological factors, leading to a complex spatio-ecological mosaic of diploid and polyploid populations. The typical ecogeographic pattern in European plants involves spatially restricted diploids growing in southern regions, interpreted as glacial refugia, and their widespread polyploid derivatives occupying larger and more northerly situated ranges with harsher environments. Whether this is true for individual ploidy-variable groups is, however, largely unknown because we lack sufficiently detailed investigations of ploidy-variable plant groups jointly applying cytological, ecological and genetic methods. We assessed ploidy and genome size variation, elevational and edaphic preferences, and plastid DNA variation within the Minuartia verna aggregate, a group of low-competitive heliophilous plants growing from the Mediterranean to Arctic Europe. Contrary to the expectations, tetraploids have a restricted distribution (Southern Europe) and inhabit a relatively narrow environmental niche. The distribution of diploids, on the other hand, spans the full range of conditions, including climatic (i.e. highest elevations and latitudes) and edaphic extremes (i.e. toxic serpentine and metalliferous substrates). The distribution pattern of the two ploidies could be explained by their distinct evolutionary histories, suggesting expansion of the diploid-dominated haplotype group accompanied by long-term persistence and local differentiation of tetraploids in refugia in the Balkan Peninsula. In summary, our study contradicts the prevailing view of polyploids as successful colonizers of novel and challenging habitats and points to the importance of combining ecological and genetic data when studying ploidy-variable species complexes.
... 1A, 1B). Notably, in a redefined circumscription of Minuartia, Dillenberger and Kadereit (2014) not only accepted Eremogone as a distinct genus, but to make it monophyletic they suggested its expansion by proposed transfer of the members of Minuartia subgen. Spergella (Fenzl) McNeill to Eremogone. ...
... The fact that the said phylogenetic analysis included two Himalayan members of subgenus Eremogoneastrum, viz., Arenaria bryophylla and A. kansuensis, both falling in the distinct Eremogone clade, strongly support the placement of Himalayan members of Eremogoneastrum in the genus Eremogone instead of Arenaria, s.s. Following these results, Dillenberger and Kadereit (2014) (Hara andTebbs 1979, Singh andDiwakar 2010). Of these, only one, A. polytrichoides, is reported from Indian western Himalaya. ...
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The genus Arenaria L. (Caryophyllaceae) in the Indian Western Himalaya is studied in detail and rearranged. Three genera, namely Dolophragma Fenzl, Eremogone Fenzl and Odontostemma Benth. ex G. Don, which were previously treated as subgenera under the genus Arenaria, are here recognized as distinct genera and corresponding species of Arenaria are transferred to them. As concluded in phylogenetic studies, subgenus Eremogoneastrum Fenzl is treated as a part of the genus Eremogone and new combinations are proposed for eight western Himalayan taxa transferred here under the genus Eremogone. Species hitherto treated in Arenaria subgenus Arenaria are retained as it is, except the sole representative of section Compressae McNeill, which is shifted under a newly described monotypic genus Himgiria. Similarly, three species hitherto under Arenaria subgenus Solitaria McNeill now form part of newly described Sino-Himalayan genus Shivparvatia. A key to the genera of Arenaria and its allies reported from Indian western Himalaya is also provided.
... The genus Arenaria L. (Alsinoideae Fenzl, Caryophyllaceae Juss.) comprises about 160 species of annual and perennial herbs mostly distributed in the northern temperate regions, the Mediterranean, and the Andes of South America (Hernández-Ledesma et al., 2015 and literature therein). The molecular data by Greenberg & Donoghue (2011), Dillenberger & Kadereit (2014), and Sadeghian et al. (2015) confirmed that Arenaria is polyphyletic. As a result, several species were included into other genera, such as Ermogone Fenzl, Honckenya Ehrh., Minuartia L., Moheringia L., Odontostemma Benth ex G. Don, Sabulina Rchb., Solitaria (McNeill) Sadeghian & Zarre, and Spergularia (Pers.) ...
... & C. Presl. (Conti et al., 2014;Dillenberger & Kadereit, 2014;Iamonico, 2014). The Flora of Chile comprises eight Arenaria species, of which three are endemic (Ulloa Ulloa et al., 2017;Rodriguez et al., 2018: 253-254;Zuloaga et al., 2019 [app. ...
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A nomenclatural study of the Arenaria names referring to the species found in Chile is presented. Three names, A. fastigiata, A. multicaulis, and A. scopulorum, are illegitimate (Arts. 53.1 and 52.1 of ICN); a new name is proposed for A. fastigiata and A. multicaulis; A. scopulorum was also nomenclaturally superfluous including the type of A. digyna (Art. 52.1 of the ICN). Arenaria digyna, A. microphylla, A. oligosperma, A. pleurantha, and A. rivularis are lectotypified, based on specimens preserved at E, HAL, K, P, and SGO (isolectotypes at B, K, P, and SGO). One epitype (at SI) was designated for A. rivularis.
... Para el tratamiento taxonómico se realizaron descripciones morfológicas de la familia, géneros y especies con base en la secuencia utilizada por , así como claves dicotómicas para su identificación. Para los nombres válidos y sinónimos se utilizaron aquellos aceptados por Dillenberger y Kadereit (2014), Pérez-Calix y Grajales-Tam (2013) y Sosa et al. (2006). Se elaboraron mapas de distribución para las especies por medio del programa QGIS ver. 2. 28.4 (QGIS, 2017), utilizando los datos tomados en campo y de los ejemplares de herbario consultados, a los ejemplares no georreferenciados se les asignaron coordenadas de acuerdo con la localidad de colecta. ...
... & Maguire son nue-vos registros para la entidad de acuerdo con la literatura consultada (García-Regalado et al., 1999;CONABIO, 2008;Pérez-Calix y Grajales-Tam, 2013;Villaseñor, 2016). Del total de especies localizadas en el estado, cinco son endémicas de México de acuerdo con Sosa et al. (2006), Pérez-Calix y Grajales-Tam (2013), Dillenberger y Kadereit (2014) y Villaseñor (2016): Cerdia virescens Moq. & Sessé ex DC., Spergularia mexicana Hemsl., Triplateia moehringioides (Moq. ...
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Antecedentes y Objetivos: La familia Caryophyllaceae tiene una amplia distribución geográfica, especialmente en regiones templadas y cálidas del hemisferio norte. Sus especies habitan principalmente en ambientes abiertos o sitios perturbados. El objetivo del presente estudio fue llevar a cabo el inventario, documentar la distribución y realizar el tratamiento taxonómico de las especies de Caryophyllaceae presentes en el estado de Aguascalientes, México. Métodos: Se realizaron colectas en el estado de Aguascalientes a partir de agosto 2012 hasta junio 2015. En cada sitio de colecta se tomaron coordenadas geográficas con base en Datum WGS 84 y se registró el tipo de vegetación. El material colectado fue identificado por medio de claves taxonómicas especializadas y cotejado con material de herbario. Para el tratamiento taxonómico se realizaron descripciones para familia, géneros y especies, así como claves dicotómicas para su identificación. Se elaboraron mapas de distribución con base en los datos tomados en campo y de los ejemplares de herbario consultados. Resultados clave: En Aguascalientes, la familia Caryophyllaceae está representada por 11 géneros y 21 especies, de las cuales cuatro son registros nuevos para el estado. La mayor cantidad de especies se encuentra en tipos de vegetación de clima templado, principalmente bosque de Quercus y bosque mixto, seguido por matorral xerófilo y pastizales. El municipio en el que se distribuye el mayor número de especies es San José de Gracia, seguido por Calvillo. Del total de especies encontradas en el estado, cinco son endémicas de México. Conclusiones: La riqueza de géneros y especies de la familia Caryophyllaceae presentes en Aguascalientes es comparable con estados de mayor tamaño. El presente trabajo amplía el conocimiento de la flora con la que cuenta el estado y de la familia Caryophyllaceae en México.
... 150 sp.), and Silene Linnaeus (1753: 416), ect. (see e.g., Kool et al. 2007, Dillenberger & Kadereit 2014, 2018, Iamonico 2013, 2018, Iamonico & Domina 2015, Iamonico 2018, Madhani et al. 2018, Sadeghian et al. 2015. ...
Article
The Flora of Turkey includes 88 Dianthus taxa. Some interesting populations belonging were observed in the Sivas province (C-Turkey). On the basis of both morphological and molecural investigations the Sivas population was here described as a new species for the science, named Dianthus hamzaoglui. A diagnosis, detailed description, distribution area, habitat, and IUCN assessment, as well as original photos were provided, as well as a comparison with the related D. burdurensis.
... Hook.f. and C. masonae L.B.Moore. For the molecular dating analyses, nine specimens from Sagina species (seven from GenBank and two newly sequenced samples) were included as outgroups, based on the relationships reported by Dillenberger and Kadereit (2014) and Greenberg and Donoghue (2011) (see Table S1 for information on samples and GenBank accession numbers). Herbarium samples were obtained from the herbaria of the British Antarctic Survey, UK, and the University of Magallanes in Punta Arenas, Chile (herbarium codes AAS and HIP, respectively). ...
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Aim Antarctica's remote and extreme terrestrial environments are inhabited by only two species of native vascular plants. We assessed genetic connectivity amongst Antarctic and South American populations of one of these species, Colobanthus quitensis, to determine its origin and age in Antarctica. Location Maritime Antarctic, sub‐Antarctic islands, South America. Taxon Antarctic pearlwort Colobanthus quitensis (Caryophyllaceae). Methods Four chloroplast markers and one nuclear marker were sequenced from 270 samples from a latitudinal transect spanning 21–68° S. Phylogeographic, population genetic and molecular dating analyses were used to assess the demographic history of C. quitensis and the age of the species in Antarctica. Results Maritime Antarctic populations consisted of two different haplotype clusters, occupying the northern and southern Maritime Antarctic. Molecular dating analyses suggested C. quitensis to be a young (<1 Ma) species, with contemporary population structure derived since the late‐Pleistocene. Main conclusions The Maritime Antarctic populations likely derived from two independent, late‐Pleistocene dispersal events. Both clusters shared haplotypes with sub‐Antarctic South Georgia, suggesting higher connectivity across the Southern Ocean than previously thought. The overall findings of multiple colonization events by a vascular plant species to Antarctica, and the recent timing of these events, are of significance with respect to future colonizations of the Antarctic Peninsula by vascular plants, particularly with predicted increases in ice‐free land in this area. This study fills a significant gap in our knowledge of the age of the contemporary Antarctic terrestrial biota. Adding to previous inferences on the other Antarctic vascular plant species (the grass Deschampsia antarctica), we suggest that both angiosperm species are likely to have arrived on a recent (late‐Pleistocene) time‐scale. While most major groups of Antarctic terrestrial biota include examples of much longer‐term Antarctic persistence, the vascular flora stands out as the first identified terrestrial group that appears to be of recent origin.
... The family Caryophyllaceae Jussieu (1789: 299) consists of ca 100 genera distributed worldwide, mainly in the northern hemisphere (Hernández-Ledesma et al. 2015). Many taxonomical and molecular studies have been carried out on the family (Bittrich 1993, Harbaugh et al. 2010, Sadeghian et al. 2015, Dillenberger & Kadereit 2014, Iamonico 2013, 2014, 2015, 2018, Iamonico & Domina 2015, Madhani et al. 2018, Koç et al. 2019 or the order Caryophyllales (Hernández-Ledesma et al. 2015). Minuartia Loefling in Linnaeus (1753: 89) belongs to Caryophyllaceae subfam. ...
Article
In this study a new species from Turkey, Minuartia alpuensis (Caryophyllaceae), is described and illustrated, and its taxonomic relationships are discussed. The distribution of the new species, its ecology and conservation status are given, and an identification key is provided. Pollen structures of both new and related taxa are examined.
... Dillenb. & Kadereit (Dillenberger and Kadereit 2014); we will hereafter refer to the species using its common name, Geocarpon. Geocarpon is extremely diminutive; mature plants reach 1-4 cm or less in height and have one to several small stems that each produce a few small pairs of leaves measuring < 4 mm in width ( Fig. 1a, b, and d). ...
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Conservation genetics studies not only provide information about genetic diversity and genetic structure to inform conservation strategies, they can also help infer life history characteristics such as mating system, pollinator, and seed dispersal strategy of a plant species. Here, we investigated Geocarpon (Mononeuria minima; Caryophyllaceae), an extremely diminutive, poorly known plant species from the south-central U.S. that is threatened due to high habitat specificity and habitat loss. The goals of this study were to use genetic data to help understand the basic attributes of the biology of Geocarpon, including mating system and the spatial extent of gene flow, how genetic variation is partitioned within and among populations and across the landscape, and how to protect that genetic variation. Most Geocarpon populations are highly homozygous and genetically homogenous, indicating a predominantly selfing mating system. Although the species maintains some allelic diversity, the majority of genetic variation was partitioned among populations, even in groups separated by small geographic distances (≥ 0.5 km), indicating very localized seed dispersal (gravity or water). Because genetic variation is structured at a fine scale, to conserve the full range of genetic diversity it is necessary to protect a large proportion of the populations of the species; we recommend protection of as many currently unprotected sites as possible, particularly in areas of the geographic range where few populations are protected, combined with ex situ conservation seed banking in sites that cannot be protected. This study illustrates how life history characteristics, particularly mating system, strongly influence patterns of genetic structure and can have major effects on the strategy to conserve genetic variation in an endangered species.
... Caryophyllaceae is a large mainly Holarctic family of approximately 3000 species of herbs and subshrubs, with its diversity center in the Mediterranean and the adjacent family have shown that many of the traditionally defined genera are not monophyletic (Dillenberger & Kadereit, 2014;Pirani et al., 2014;Sadeghian et al., 2015;Madhani et al., 2018). This is also the case for the tribe Caryophylleae, where, based on morphology (connate sepals, stipitate ovary, and presence of two styles), the plant from the Shirkuh Mts. was suspected to belong to. ...
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Although mountain ranges are often recognized as global biodiversity hotspots with a high level of endemism, diversity and biogeographic connections of isolated and weakly explored mountains remain poorly understood. This is also the case for Shirkuh Mts. in central Iran. Here, Yazdana shirkuhensis gen. & spec. nov. (Caryophylleae, Caryophyllaceae) is described and illustrated from the high alpine zone of this mountain. Molecular phylogenetic analyses of nuclear and plastid DNA sequence data show that Yazdana shirkuhensis is related to Cyathophylla and Heterochroa (tribe Caryophylleae). The herein newly described genus and species accentuate the Shirkuh Mts. as a center of endemism, which harbors a high number of narrowly distributed species, mostly in high elevations reaching alpine habitats. As this area is currently not protected, high elevations of Shirkuh Mts. have conservation priority. This article is protected by copyright. All rights reserved.
... proposed by Harbaugh et al. (2010), Greenberg & Donoghue (2011), Dillenberger & Kadereit (2014), Sadeghian et al. (2015), and Pusalkar & Singh (2015). In the revised circumscription Arenaria s.l. is divided into five genera i.e. ...
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Information about trichomes diversity and distribution of the family Caryophyllaceae is rare and the present work is intended to fill this knowledge lacuna. In the present work 62 taxa belonging to 19 genera were studied. For the analysis of trichomes diversity and vestiture type, dried plant specimens were rehydrated with water. The final illustrations of trichomes were made by using camera lucida. Six types of trichomes viz., Unicellular eglandular, Unicellular glandular, Multicellular uniseriate glandular, Multicellular uniseriate eglandular, Multicellular eglandular bifurcate and Multicellular multiseriate eglandular trichomes reported in the studied taxa. Diversity of trichome and their distribution does not play any significant role in the taxonomic delimitation either generic or tribal level of the family Caryophyllaceae. Although, few closely allied species can be distinguished from each other either on the basis of the presence of trichomes or vestiture patterns.
... Οι τρεις παραδοσιακές υποοικογένειες Silenoideae, Alsinoideae και Paronychioideae (Pax, 1899;Bittrich, 1993) δεν φαίνεται να επιβεβαιώνονται από τα ευρήματα της μοριακής φυλογενετικής, επομένως έχουν εγκαταληφθεί στις πιο πρόσφατες θεωρήσεις της οικογένειας, αντικαθιστώμενες από 11 φυλές (tribes) συγγενικών γενών (Ηarbaugh et al., 2010;Greenberg & Donoghue, 2011). Αντιστοίχως, σε επίπεδο γενών αναμένονται αρκετές ανακατάταξεις τα επόμενα χρόνια χάρη στην ευρύτερη και πιο ευχερή χρήση των μοριακών δεικτών, με χαρακτηριστικό παράδειγμα τη διάσπαση του πολυφυλετικού γένους Minuartia σε 11 γένη (Dillenberger & Kadereit, 2014). ...
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Dianthus cruentus Griseb. (Caryophyllaceae) is a herbaceous perennial species native to the Balkan Peninsula, including Greece, and western Asia. It is a species of great ornamental interest due to its architectural inflorescences and its compact blue-green foliage, particularly resistant to dry and poor soils and attractive to pollinators such as butterflies and bees. In addition, it is a species of medical potential due to its strong antioxidant action. In the present study, the determination of the appropriate conditions for maximizing the germination of its seeds in both in vitro and in vivo conditions was initially investigated. Two seed lots collected from a native population of the species on Mount Kallidromo, Fthiotida, dry-stored for 0 or 12 months, were used, without any pre-treatment. Regarding in vitro germination, the seeds were surface sterilized with a 20% commercial bleach water solution for 10 minutes, were then rinsed with distilled water and afterwards were placed in Petri dishes containing ½ MS medium, under 16 h photoperiod from 37.5 μmol m-2 s-1 fluorescent light, at temperatures of 5, 10, 15, 20 or 25 °C. The optimum temperature for seed germination was 15 °C, in which the highest germination ability in both recently collected (98%) and 12-month-old seeds (100 %) was observed. The lowest germination percentages were recorded at the two extreme temperatures of 5 °C and 25 °C, with particularly low values for recently collected seeds (18% and 15%). At all temperatures, minus that of 5 °C, the seeds germinated rapidly (T50 = 2-6 d). The duration of the seeds’ dry storage significantly affected their germination, increasing the germination percentage at the 2 extreme temperatures and demonstrating the positive effect of dry afterripening. As for ex vitro germination, the seeds were placed in pots with a mixture of peat-perlite (1: 1, v/v) and were incubated in growing chambers at temperatures of 15, 20 or 25 °C. The highest germination percentage was observed at 15 °C for both seed lots (97-99%), while the lowest at the temperature of 25 °C (62% and 4% for the 0- and 12-months-old seeds respectively). Germination ability was slightly lower in ex vitro germination, especially at 25 °C. Furthermore, the micropropagation of the species from explants excised from 2-month-old seedlings was investigated on solid substrates. Substrates containing MS medium and 8 g L-1 agar were used in almost all substages of the proliferation stage, and the effects of cytokinins BA, 2iP and ZEA, with the absence or the presence of auxin NAA, were investigated, using explants derived from non-hyperhydric or hyperhydric shoots. In the initial culture substrates containing 0.1 mg L-1 BA or 0.1 or 0.5 mg L-1 2iP the explants produced shoots at a high percentage (72-87%), but many of these were shown to be hyperhydric, with abnormal morphology and vitreous appearance. The number of hyperhydric shoots and the number of nodes seemed to be higher on substrates containing 2iP, with the largest number of normal shoots (2.3) appearing in the substrate containing 0.1 mg L-1 BA. During the 1st subculture stage, hyperhydric shoots were observed in bigger numbers in the substrate containing 2iP, while an increase in shoot proliferation and the number of hyperhydric shoots was correlated with the use of hyperhydric explants grown on hormone-free MS substrate. In the 2nd subculture stage, the presence of 0.1 mg L-1 BA yielded the largest number of shoots regardless of the physiology of the explants, while the addition of 0.05 mg L-1 resulted in a significant increase in the number of normal shoots and a reduction in hyperhydricity rates when hyperhydric explants were used. In the 3rd subculture, when explants derived from nonhyperhydric shoots were used the highest value of the multiplication index (4.5) and the maximum number of shoots (3.6) was recorded in MS substrate containing 2 mg L-1 BA. Increasing the concentration of agar to 12 g L-1 produced the largest number of shoots (2.4) when hyperhydric explants were used. In both cases, the highest percentages of hyperhydricity were recorded on MS substrate containing 0.1 mg L-1 ZEA. In contrast, during the 4th subculture, the use of this substrate resulted in the greatest multiplication index value at this stage (4.8). Overall, the use of MS substrate with 0.1 mg L-1 BA and 0.05 mg L-1 NAA resulted in the highest values of the proliferation index (5.1) regardless of the physiology of the explants used, with its maximum value (5.2) recorded from the culture of hyperhydric explants on MS substrate containing 0.1 mg L-1 BA and 12 g L-1 agar. The presence of phytohormones in the substrates increased the production of callus around the base of the explants, while high concentrations of cytokinins lessened the occurence of roots. D. cruentus microshoots were rooted in MS or ½ MS substrates in the presence or absence of IBA. The use of MS substrate resulted in lower rooting rates (55-70%) when lower concentrations of IBA were used (0 or 0.1 mg L-1). The optimal concentration of IBA was found to be 0.5 mg L-1 regardless of the multiplication substrate where the microshoots originated or the nutrient medium used in the rooting substrate, with rooting rates of 83-100%, while in some instances a carry over effect, caused by the cytokinin content of the shoot proliferation substrates, was observed on the roots’ number and median length. The ex vitro acclimatization of the rooted microshoots was completed with a success rate of 91% using a peat:perlite substrate (1:1, v/v). Significant morphological differences were observed between acclimatized plantlets, including differences in stem length and total number of shoots and nodes, which were correlated with the influence of different shoot proliferation and/or rooting substrates, while the presence of IBA during rooting seemed to have an amendable effect on their survival. Finally, the use of an integrated, 60-days-long propagation protocol with shoot proliferation and rooting occurring concurrently in one stage on MS substrates free of PGRs or containing BA or 2iP and NAA produced rooted young plants at a high percentage (88%), which had the maximum observed total number of shoots and nodes and were then successfully acclimatized, surviving at a percentage of 86%. Scientific area: Horticulture Keywords: Dianthus cruentus, native ornamental plant, germination ecophysiology, seed germination in vitro, seed germination temperature, seed lot age, dry afterripening, in vitro propagation, solid culture, seedling explant, hyperhydricity, hyperhydric explant, callus induction, in vitro rooting, ex vitro acclimatization, cytokinins, IBA, carry over effect, acclimatized plantlets’ morphology
... aus Minuartia s. l. ausgegliedert werden. Dillenberger & Kadereit (2014) konnten zeigen, dass die verwendeten Merkmale zur Gattungsabgrenzung von Minuartia völlig ungeeignet waren. Die mehrfache Ausschlüsselung von Minuartia im Hauptschlüssel der Familie hat schon in früheren Auflagen des Grundbandes gezeigt, dass Minuartia s. l. morphologisch divers ist und sich schwer zusammenfassen lässt, jedoch die Aufteilung in vier natürliche Gattungen erleichtert. ...
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The 22nd edition of the Rothmaler field flora was published in 2021, and the present article gives comments on the numerous, especial major changes. We added new species, and for the first time included an Algae group; the stoneworts, Characeae, which in many practical respects resemble vascular plants. For all species, biological data were revised, and chromosome numbers were added if available for accessions from Germany. Notes on distribution within Germany were completely restructured, employing new physico-geographical rather than political mapping units, and using data from the current distribution atlas of vascular plants in Germany. Taxonomy and nomenclature in the Rothmaler largely follow the standard checklist of vascular plants in Germany; deviations are explained and justified. Die 22. Auflage des Gundbandes der Rothmaler Exkursionsflora ist 2021 erschienen und der vorliegende Artikel gibt Kommentare und Hinweise zu vielen, vorwiegend größeren Neuerungen. Wir haben im Grundband nicht nur neue Taxa aufgenommen, sondern mit den Armleuchteralgen, den Characeae, auch erstmals eine Algengruppe verschlüsselt, die aber in vieler, insbesondere ökologisch-praktischer Hinsicht durchaus den Gefäßpflanzen ähnelt. Für alle Arten im Band wurden die biologischen Angaben durchgesehen; sofern Daten zu Akzessionen aus Deutschland verfügbar waren, wurde Chromosomenzahlen neu eingefügt. Die Angaben zur Verbreitung in Deutschland wurden basierend auf dem Deutschlandatlas neu zusammengestellt, wobei jetzt stärker naturräumliche als politische Verbreitungseinheiten genutzt wurden. Taxonomie und Nomenklatur im Grundband folgen weitgehend der Florenliste von Deutschland, Abweichungen werden hier diskutiert und ggf. begründet.
... or Eken et al. (2006), where area-based lists of selected rare species are offered. The naming of the species follows largelyGüner et al. (2012), with the exception of the following taxa (references to the taxonomic backbones in brackets): Anemonastrum, Asteraceae, Lomelosia, Noccaea, Odontarrhena and pteridophytes (The Euro+Med PlantBase [http://ww2.bgbm.org/EuroPlusMed/query. asp]; Caryophyllales(Dillenberger and Kadereit 2014;Hernández-Ledesma et al. 2015;Madhani et al. 2018), Lamium ...
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Abstract. This chapter characterizes the flora and vegetation of the North-Western Caucasus, taking the Karachaevo-Cherkessian Republic (a mountainous region in the southwestern Russia) and the Teberda State Biosphere Reserve (a protected nature area with restricted access located on the northern spurs of the Greater Caucasus Range) as examples. The Teberda State Biosphere Reserve is located in the southern part of the Karachaevo-Cherkessian Republic at the upper reaches of the Teberda and Kizgych rivers and their tributaries. The elevation in the Teberda State Biosphere Reserve varies from 1259 to 4046 m a.s.l. The treeline runs at 1700–2400 m a.s.l., and the upper limit of vascular plants is found at 3750 m a.s.l. About 83% of the Reserve area lies above 2000 m a.s.l. Rock and scree outcrops occupy 26% of the Reserve. The wide elevational range, complex topography, and sharp changes of microclimatic conditions determine a high species diversity and variety of plant communities. The flora of the Karachaevo-Cherkessian Republic currently comprises 1959 species of vascular plants. There are 10 floristic elements in the indigenous flora of the Republic, i.e. Holarctic, Palaearctic, Panboreal, Euro-Mediterranean, Mediterranean, Ancient Mediterranean, Irano-Turanian, Pontic-South- Siberian, Pluriregional, and Endemic. A total of 343 species (17.5%) are endemics. Six elevational belts (steppe, deciduous broad-leaved forest, coniferous forest, subalpine, alpine, and subnival) and major vegetation types, including steppes, deciduous and coniferous forests, crooked-stem elfin woods, tall grass communities, fens, alpine and subalpine meadows, snowbeds, lichen heaths, pioneer communities on rocks, screes and talus slopes are reviewed. The Caucasus is considered a biodiversity hotspot with significant levels of endemism at both species and ecosystem levels. Currently it is prioritized as one of the most vulnerable regions with a high irreplaceability.
... Given the dynamic range history of plant taxa in the EAS, we here examine the role of hybridization in the evolution of a subclade of Cherleria L. (formerly Minuartia L., Caryophyllaceae; [21,54]) endemic to the EAS (Fig. 1). Cherleria as a whole contains 19 species and has a circumboreal distribution, with incursions into the high mountains of Eurasia and North America. ...
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Background Cherleria (Caryophyllaceae) is a circumboreal genus that also occurs in the high mountains of the northern hemisphere. In this study, we focus on a clade that diversified in the European High Mountains, which was identified using nuclear ribosomal (nrDNA) sequence data in a previous study. With the nrDNA data, all but one species was monophyletic, with little sequence variation within most species. Here, we use genotyping by sequencing (GBS) data to determine whether the nrDNA data showed the full picture of the evolution in the genomes of these species. Results The overall relationships found with the GBS data were congruent with those from the nrDNA study. Most of the species were still monophyletic and many of the same subclades were recovered, including a clade of three narrow endemic species from Greece and a clade of largely calcifuge species. The GBS data provided additional resolution within the two species with the best sampling, C. langii and C. laricifolia , with structure that was congruent with geography. In addition, the GBS data showed significant hybridization between several species, including species whose ranges did not currently overlap. Conclusions The hybridization led us to hypothesize that lineages came in contact on the Balkan Peninsula after they diverged, even when those lineages are no longer present on the Balkan Peninsula. Hybridization may also have helped lineages expand their niches to colonize new substrates and different areas. Not only do genome-wide data provide increased phylogenetic resolution of difficult nodes, they also give evidence for a more complex evolutionary history than what can be depicted by a simple, branching phylogeny.
... Molecular studies on Petrorhagia, Velezia and Gypsophila revealed the relationship among related taxa (Korkmaz & Doğan 2015;Poyraz et al. 2012;Hilooğlu et al. 2016). Molecular phylogenetic studies based on ITS and plastid trnQ-rps16 DNA in the largest genera (Gypsophila L., Dianthus L., Arenaria L., Minuartia L. and Silene L.) have resulted in revised treatments including new genera and synonymy (Dillenberger & Kadereit 2014;Madhani et al. 2018;Sadeghian et al. 2015). ...
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Gypsophila malyerii Hamzaoğlu & Koç, a new species of sect. Capituliformes, is described and illustrated from Turkey. Information on distribution, habitat and conservation status are given. The most similar species is G. osmangaziensis. The morphology and micromorphology of seed and pollen characters of the two closely related species are compared.
... Augustea Iamonico, Facchinia Rchb., Mcneillia Dillenb. & Kadereit, Sagina L., Scleranthus L.;Smissen et al., 2003;Dillenberger & Kadereit, 2014;Iamonico, 2015Iamonico, , 2016 y, en algunos casos, merecen aclaraciones sobre su nomenclatura y taxonomía. Tal es el caso de Cardionema DC., un género nativo del Nuevo Mundo, con unas 6 especies, distribuido desde los Estados Unidos de América hasta el sur de la Patagonia (Hartman, 2005;Ulloa Ulloa et al., 2017;Hernández-Ledesma et al., 2015). ...
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The genus Cardionema (Caryophyllaceae) is native to the New World and it comprises six species of perennial herbs, with spiny leaves and sepals, membranous stipules and bracts, and reduced petals. This study aims to update the richness, morphology and distribution of Cardionema in Argentina, based on the study of herbarium specimens. Four species of Cardionema are accepted for the Flora of Argentina; a description of each taxon, an identification key, illustrations and pictures, a comparative-morphology based table, and distribution maps are included. Two lectotypes are also designated.
... 3000 species, mainly distributed in the northern hemisphere, Mediterranean and Irano-Turanean regions (Hernandez-ledesma et al. 2015). Several Caryophyllaceae genera are critical from the taxonomical and nomenclatural point of view (see e.g., (Aydın et al. 2014, Dillenberger & kadereit 2014, Iamonico & Domina 2015, Fedoronchuk & Mosyakin 2016, Iamonico 2016) and need still studies. Among these genera, Saponaria linnaeus (1753: 408). ...
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Saponaria karapinarensis (Caryophyllaceae), an endemic species from Turkey, was studied from taxonomic point of view. Morphological, anatomical, and palynological characteristics of the species were examined. Morphology of stem, calyx, petals, and capsule are useful to distinguish the species. Anatomical features, such as the number of cortex layers and sclerenchymatic cell layers in stem, are of taxonomical importance. SEM Scanning Electron Microscopy (SEM) studies on the pollen grains have revealed that they are spheroideae and their exine ornamentation is granulatae, spinulate microechinatae- microperforatae. Seed micromorphological features were also given.
... & J.Presl including about 3000 species (100 genera) which are primarily distributed in the Holarctic region, having a center of diversity in the Mediterranean and Irano-Turanian regions (Bittrich 1993, Heywood 1998, Hernández-Ledesma et al. 2015. Greenberg & Donoghue (2011) demonstrated the monophyly of Caryophyllaceae using molecular data, but its classification remains partially unresolved since several genera are critical from the taxonomical and nomenclatural point of view (Conti et al. 2014, Dillenberger & Kadereit 2014, Iamonico 2015, 2021a, 2021b, Iamonico & Domina 2015. ...
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The seed and leaf structures of some species belonging to the genus Saponaria (Caryophyllaceae) were examined using SEM. These species were: S. calabrica, S. dalmasii (endemic), S. glutinosa, S. halophila (endemic), S. karapinarensis (endemic), S. kotschyi (endemic), S. mesogitana, S. officinalis, S. orientalis, S. pamphylica (endemic), S. picta (endemic), S. pinetorum var. elatior (endemic), S. prostrata subsp. anatolica (endemic), S. prostrata subsp. calvertii, S. prostrata subsp. prostrata (endemic), S. pumilio, and S. tridentata. The results obtained show that the micromorphology of the seed are useful to distinguish the studied species. Four groups can be distinguished using the number of seed per capsule. In addition, the shape of the seed, microsculpture, shape and margin of testa cells were found as important characteristics to identify the various species. Leaf anatomy reveals that trichome type and mesophyll symmetry define significant differences between the studied species.
... Six out of these nine latter species were later recognized as part of the following two other genera: Mononeuria Rchb. Dillenberger and Kadereit, 2014) which is sister to some Minuartia members. They need further study and are part of an ongoing study by the author of the present paper (Iamonico, in prep.). ...
Article
Background and Aims: Stellaria traditionally comprises 150-200 species, mainly distributed in the temperate regions of Eurasia and North America. Molecular studies demonstrated that Stellaria is polyphyletic and includes about 120 species. The genus has a high phenotypic variability which has led to nomenclatural disorders, making the identification of the various species difficult. A note is presented about a taxon currently accepted under the genus Stellaria -Stellaria obtusa- which should be recognized as a separate genus, here proposed as Engellaria gen. nov. Methods: This study is based on examination of specimens of American and European herbaria and analysis of relevant literature. Key results: Available molecular data show that Stellaria obtusa is not included in the Stellaria s.s. clade, but instead is basal to another clade comprising the genera Honckenya, Schiedea, and Wilhelmsia. Stellaria obtusa was, therefore, compared with these three groups and with morphologically similar apetalous members of Stellaria s.s. (S. crispa, S. media, S. pallida, and S. irrigua). The results obtained lead to the recognition of S. obtusa as a separate new North American monotypic genus. A diagnostic key of the apetalous members belonging to the American Caryophyllaceae genera is proposed. Finally, the names Stellaria obtusa and S. washingtoniana (= S. obtusa) are lectotypified based on specimens deposited, respectively, at UC (isolectotypes at GH, NY, and YU) and GH (isolectotypes at BM, CAN, CAS, CS, DOV, F, GH, K, MIN, MSC, NY, US, and VT). For the name Alsine viridula (= S. obtusa) the holotype was found at US (isotypes at CAS, F, GH, NY, OSC, RM, and UC). Conclusions: Stellaria obtusa does not belong to the genus Stellaria. The present study shows that the combined use of morphological data and phylogenetic analyses helped to clarify the taxonomic position of difficult plant groups, as in Stellaria.
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Whereas the eastern North American–eastern Asian floristic connection represents one of the most widely studied biogeographical relationships in flowering plant evolution, connections between western North America and Asia have been comparatively rarely investigated, especially through genetic approaches. Stellaria irrigua is one of several plants that has been treated as an exceptionally dramatic example of a disjunction between floristically similar, high alpine biotas of the southern Rocky Mountains and south‐central Siberia. We here employ numerous new field collections and ddRADseq data to test the hypothesis that S. irrigua—a species that has been known for over 180 years—represents a long‐distance disjunction between the southern Rocky Mountains and central Asia. Extensive fieldwork, review and perusal of herbarium materials, and phylogenomic analyses indicate that S. irrigua is broadly distributed across an amphi‐Beringian arc extending from southern and central Asia, east through Beringia, and south throughout mountainous regions of western North America. Sampled Asian populations formed two clades, and North American individuals all formed a clade embedded within this broader Asian lineage. Stellaria irrigua is, however, rendered non‐monophyletic by a lineage that is embedded within the North American populations and is ecologically and morphologically distinctive from S. irrigua. The identity of this newly recognized lineage, which was in prior works attributed to S. irrigua, has been confused since plants of the former were first collected in the San Juan Mountains of Colorado in the late 1800s under Arenaria and Alsine. We provide a new name for this taxon, Stellaria sanjuanensis, a charismatic starwort of dry alpine scree slopes of the southern Rocky Mountains. Additionally, two lectotypes are designated, one holotype and one isotype are identified, and two new synonymies are proposed, to help stabilize the taxonomy and nomenclature of this long‐confused species complex. A key to the starworts of the southern Rocky Mountains is also provided, and Stellaria alsine is reported as new to the region.
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The new combination Myosotis paucipilosa is validated, and differences between Myosotis refracta s.str. and M. paucipilosa are given.
Chapter
This contribution provides a survey on the high-mountain vegetation of Anatolia, Turkey, covering the West Anatolian Mts., the Taurus mountain system, the Inner Anatolian volcanoes, the East Anatolian highlands and the Black Sea Mts. Due to its location between the Black Sea and the Mediterranean Sea, the intersection of three phytogeographical regions, a highly varied geologic and climatic setting in addition to a dramatic geological past, the plant life of the different Anatolian mountains systems and isolated peaks is amazingly diverse and very rich in endemics. The chapter introduces all important high-mountain ranges, their zonation, major ecosystems, key vegetation types and floristic inventories. Based on a thorough phytogeographic analysis of the high-mountain flora and an evaluation of a wide range of floristic and phylogenetic studies, the diversity patterns of the Anatolian mountain systems and their floristic links to the adjacent areas are reviewed and mapped.
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Minuartia smejkalii is an obligate serpentinophyte plant endemic to the Czech Republic. Since the 1960s, the species’ habitat has undergone strong human-mediated fragmentation, resulting in extinction of some populations and dramatic size reduction of the remaining populations. Thus, contrary to the typically stable serpentine habitats, M. smejkalii habitats underwent a recent and severe decline, which can exacerbate the effects of fragmentation on population genetic structure. We examined the genetic structure of all known M. smejkalii populations and two populations of M. corcontica and M. caespitosa, which are closely related, using RADSeq. The results indicate low, but clear differentiation among the three species, thus supporting the status of M. smejkalii as an independent taxon, though more extensive analysis of the whole group is needed. We further show high genetic diversity within M. smejkalii populations, low to moderate among-populations differentiation, and moderate regional differentiation. This could be due to the outcrossing mating system of M. smejkalii promoting high levels of gene flow and historical factors (multiple founder events, a recent bottleneck and/or a genetic time lag). We finally demonstrate that 2–3% of the markers show differentiation patterns consistent with divergent selection, suggesting that some local adaptation might have occured in M. smejkalii. Based on our observations, but without any experimental testing for local adaptation, if a conservation action is to be carried out, we recommend strictly separating the material from the two regions, and if possible, separating the populations within a region.
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The present edition of the annotated checklist is a comprehensive catalogue of all vascular plant taxa: native and alien that occur in Iceland. The checklist features nearly 2500 taxa names, including ca. 1000 accepted names and more than 1400 synonyms and encompasses, apart from the updated list of native taxa, a complete and revised list of non-native plants (both naturalized and casual) as well as a number of more important cultivated species. According to the checklist, there are 426 native taxa in the Icelandic flora. Ten taxa have been classified as doubtfully native, ten taxa have been classified as non-native of unknown age and 19 taxa qualified as archaeophytes. There are at least 65 non-native taxa naturalized in the Icelandic flora. In total, there are 530 taxa able to form self-sustaining populations in Iceland. Apart from the main core, 282 taxa have been registered as casual aliens (not able to form self-sustaining populations). One species – Primula egaliksensis, has been classified as extinct. The list encompasses also 150 taxa excluded from the Icelandic flora, with brief explanations of the reasons that lead to the exclusion.
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Background Cherleria (Caryophyllaceae) is a circumboreal genus that also occurs in the high mountains of the northern hemisphere. In this study, we focus on a clade that diversified in the European High Mountains, which was identified using nuclear ribosomal (nrDNA) sequence data in a previous study. With the nrDNA data, all but one species was monophyletic, with little sequence variation within most species. Here, we use genotyping by sequencing (GBS) data to determine whether the nrDNA data showed the full picture of the evolution in the genomes of these species. Results The overall relationships found with the GBS data were congruent with those from the nrDNA study. Most of the species were still monophyletic and many of the same subclades were recovered, including a clade of three narrow endemic species from Greece and a clade of largely calcifuge species. The GBS data provided additional resolution within the two species with the best sampling, C. langii and C. laricifolia, with structure that was congruent with geography. In addition, the GBS data showed significant hybridization between several species, including species whose ranges did not currently overlap. Conclusions The hybridization led us to hypothesize that lineages came in contact on the Balkan Peninsula after they diverged, even when those lineages are no longer present on the Balkan Peninsula. Hybridization may also have helped lineages expand their niches to colonize new substrates and different areas. Not only do genome-wide data provide increased phylogenetic resolution of difficult nodes, they also give evidence for a more complex evolutionary history than what can be depicted by a simple, branching phylogeny.
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The complete plastome sequences of six species were sequenced to better understand the evolutionary relationships and mutation patterns in the chloroplast genome of the genus Colobanthus. the length of the chloroplast genome sequences of C. acicularis, C. affinis, C. lycopodioides, C. nivicola, C. pulvinatus and C. subulatus ranged from 151,050 to 151,462 bp. The quadripartite circular structure of these genome sequences has the same overall organization and gene content with 73 protein-coding genes, 30 tRNA genes, four rRNA genes and five conserved chloroplast open reading frames. A total of 153 repeat sequences were revealed. Forward repeats were dominant, whereas complementary repeats were found only in C. pulvinatus. the mononucleotide SSRs composed of A/t units were most common, and hexanucleotide SSRs were detected least often. eleven highly variable regions which could be utilized as potential markers for phylogeny reconstruction, species identification or phylogeography were identified within Colobanthus chloroplast genomes. Seventy-three protein-coding genes were used in phylogenetic analyses. Reconstructed phylogeny was consistent with the systematic position of the studied species, and the representatives of the same genus were grouped in one clade. All studied Colobanthus species formed a single group and C. lycopodioides was least similar to the remaining species.
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Background: Cherleria (Caryophyllaceae) is a circumboreal genus that also occurs in the high mountains of the northern hemisphere. In this study, we focus on a clade that diversified in the European High Mountains, which was identified using nuclear ribosomal (nrDNA) sequence data in a previous study. With the nrDNA data, all but one species was monophyletic, with little sequence variation within most species. Here, we use genotyping by sequencing (GBS) data to determine whether the nrDNA data showed the full picture of the evolution in the genomes of these species. Results: The overall relationships found with the GBS data were congruent with those from the nrDNA study. Most of the species were still monophyletic and many of the same subclades were recovered, including a clade of three narrow endemic species from Greece and a clade of largely calcifuge species. The GBS data provided additional resolution within the two species with the best sampling, C. langii and C. laricifolia, with structure that was congruent with geography. In addition, the GBS data showed significant hybridization between several species, including species whose ranges did not currently overlap. Conclusions: The hybridization led us to hypothesize that lineages came in contact on the Balkan Peninsula after they diverged, even when those lineages are no longer present on the Balkan Peninsula. Hybridization may also have helped lineages expand their niches to colonize new substrates and different areas. Not only do genome-wide data provide increased phylogenetic resolution of difficult nodes, they also give evidence for a more complex evolutionary history than what can be depicted by a simple, branching phylogeny.
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Establishing a baseline of current Arctic vascular plant diversity and distribution is critical, given the rapid and major environmental changes occurring in the Arctic ecozone in response to climate change. Here, we report the results of a floristic study of vascular plant diversity of Dorset and Mallik islands in the Canadian Arctic Archipelago, Nunavut, Canada. These two small islands lie off the coast of the Foxe Peninsula of southwestern Baffin Island, and they are part of the Circumpolar Arctic bioclimate Subzone C. The hamlet of Kinngait (formerly Cape Dorset) is located on Dorset Island, and Nunavut’s Mallikjuak Territorial Park encompasses all of Mallik Island. The study is based on a specimen-based dataset comprising 876 unique collections from the two islands gathered over the last century, including 268 new ones collected in 2015. Results are presented in an annotated checklist. The vascular plant flora of the study area comprises 26 families, 71 genera, 150 species and three infraspecific taxa; 139 species are recorded on Dorset Island and 102 on Mallik Island. Eleven taxa are newly recorded from the study area in six families: Carex rupestris, Eriophorum scheuchzeri subsp. scheuchzeri, E. triste (Cyperaceae); Diapensia lapponica (Diapensiaceae); Equisetum arvense subsp. alpestre (Equisetaceae); Oxytropis deflexa var. foliolosa (Fabaceae); Potentilla arenosa subsp. arenosa, P. hyparctica subsp. hyparctica (Rosaceae); Antennaria friesiana subsp. friesiana, Askellia pygmaea, and Taraxacum phymatocarpum (Asteraceae).
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Pseudocerastium is a monotypic genus in Caryophyllaceae endemic to China. The genus has been widely accepted since it was described in 1998, however its phylogenetic position within Caryophyllaceae has never been studied. In the present study, the whole plastid genome and nuclear ribosomal internal transcribed spacer (ITS) sequences of P. stellarioides was obtained through genome skimming, and the phylogenetic position of the species was studied for the first time. Plastid phylogenomic analysis of Caryophyllaceae revealed that Pseudocerastium is clustered within the tribe Alsineae with strong support. Phylogenetic analyses based on an enlarged taxon sampling of Alsineae using five DNA regions (matK, rbcL, rps16 intron, trnL-F and ITS) revealed that P. stellarioides was nested deeply within Cerastium with strong support. Analyses of morphological character evolution suggest that the ancestral states in Alsineae include three styles and a six-lobed capsule at the apex, while both Cerastium and Pseudocerastium have five styles and ten lobes at the apex of the capsule, further supporting their close relationship. The species P. stellarioides is similar to C. wilsonii in morphology, but differs in having villous indumentum on the lower part of the filaments and compressed globose seeds. Therefore, based on the present molecular and morphological evidence, the generic name Pseudocerastium is reduced here as a new synonym of Cerastium and the species P. stellarioides is transferred to Cerastium as C. jiuhuashanense.
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The taxonomic identity and phylogenetic relationships of several South African perennial taxa often synonymised to the European Spergularia media are discussed. In particular, the case of Arenaria glandulosa Jacq., a species described as native to the Cape region of South Africa, was revisited. We found this taxon to be a Spergularia, endemic to salt-laden coastal (and rarely also saline inland) habitats of the Western Cape and Eastern Cape provinces of South Africa. This taxon is a prostrate to procumbent caespitose herb, densely glandulous all over, with small flowers solitary at the base of each leaf-like bract or in congested dichasial cymes, producing petals about equalling the sepals and free styles, among other characters. Further, we describe a morphologically similar species (often mistaken for S. media and S. bocconei) occurring in succulent Karoo scrub of Namaqualand, as a new species, named S. namaquensis. Both are glandulous dwarf subshrubs considerably differing from the recently discovered S. quartzicola, a soil-specialist endemic from Namaqualand quartzitic outcrops, by many notable vegetative and reproductive features. Molecular analyses of plastid (trnL–trnF region) and nuclear ribosomal (5.8S-ITS2 region) DNA sequence data support the morphological differentiation between the South African species S. glandulosa, S. namaquensis and S. quartzicola, representing an independent lineage when compared with the Northern Hemisphere members of the S. maritima group. Synonymy, new complete descriptions and ecological and distributional data are provided to facilitate further identification of species within the putative S. media group.
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Dianthus berkayii belongs to sect. Fimbriati was described and illustrated as a new species from Erzincan Province in Eastern Anatolia, Turkey. In the current study morphological, palynological, and molecular characters were compared and discussed with its closely related species. The new species is similar to D. crinitus with fimbriate corolla, linear, acuminate, ± rigid leaves. Also similar to D. vanensis with fimbriate corolla, linear, acuminate, ± rigid, curved divaricate leaves. But distinctly different from D. crinitus by having linear and curved divaricate leaves, stem with 3–4(–5) internodes (not 5–8), a style much longer than petals, and different from D. vanensis with narrower leaves, ebarbulate and long fimbriate corolla. The bracts of D. berkayii are 1/3 of the length of the calyx, but not equal to the calyx length. The IUCN threat category of D. berkayii was determined to be “CR (Critically Endangered)” therewithal it is an Irano-Turanian element. In the study, the geographical distributions of D. berkayii, D. crinitus, and D. vanensis are also mapped, the identification key of these species was provided, pollen morphology and phylogenetic analyses based on ITS region of rDNA and ecological notes are also presented.
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Generic delimitation in Caryophyllaceae has been challenging and has been informed most recently by use of molecular phylogenetic data. In this study, analysis of 29 samples from the small segregate Mononeuria using nuclear ITS and plastid rps16 data revealed it to be polyphyletic. The type species, (M. patula) and two other species (M. muscorum and M. paludicola) were shown to belong to the Sabulina clade. The remaining species formed a clade that also included the previously monotypic Geocarpon and was sister to a heterogeneous group that included the Hawaiian Schiedea and three other monotypic genera, Honckenya, Wilhelmsia and Triplateia. Although several nomenclatural options are available, we propose to place the species from this clade into a single genus, Geocarpon, which basically follows the most recent treatment after exclusion of Sabulina spp., but with the necessary new genus placements. New combinations are proposed: Sabulina muscorum; Sabulina paludicola; Geocarpon carolinianum; Geocarpon cumberlandensis; Geocarpon glabrum; Geocarpon groendlandicum; Geocarpon nuttallii and Geocarpon uniflorum. Analysis of the sequence data revealed remarkable variability among populations of Sabulina patula (formerly Mononeuria patula), Sabulina paludicola (formerly Mononeuria paludicola) and Geocarpon groenlandicum (formerly Mononeuria groenlandica), suggesting that cryptic species may be present. The data also suggested that broader sampling of Sabulina and Geocarpon could lead to increased understanding of the timing and origins of occupation of calcareous glades and rock outcrop habitats in eastern North America.
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The status of the genus Dichodon has long been debated, and its taxonomic position in tribe Alsineae has been changed during the time from a section or subgenus in Cerastium to genus sister to Holosteum. This group comprises important members of wet meadows in alpine and subalpine vegetation of Europe, arctic regions, and SW-Asia plus one species known as a weed in N-America, and a further one occuring in mountains of Taiwan. In order to clarify the taxonomic questions concerning this group and its species delimitation, we constructed phylogenetic trees, selecting several species belonging to tribe Alsineae as representatives of major lineages of this tribe as well as several accessions of Dichodon. Morphological studies focused more intensively on members of Dichodon using herbarium specimens and direct field examinations. The results confirm those of recent molecular phylogenetic studies, indicating Dichodon as a monophyletic genus sister to Holosteum and not Cerastium. In addition, the obtained cladograms support five distinct groups in Dichodon corresponding to five species of this genus we recognize in Iran, the focal area of this study. Seed micromorphology provides strong support for the recognition of Dichodon as a separate genus, but it is not informative at species and subspecies ranks due to constancy of most of seed characters within the genus. As part of this study, a new species—Dichodon alborzensis—is described, D. kotschyi is reported in Iran for the first time, and Cerastium schischkinii is placed in synonymy (new synonymy) under D. kotschyi.
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Nucleotide sequences of the plastid encoded gene matK were examined for their potential utility in phylogenetic analyses within angiosperm families. Sequences 661 bases in length were obtained from twenty species of Polemoniaceae. Phylogenetic analyses resulted in four equally parsimonious trees with a consistency index of 0.70. Several well supported groups allowed us to test hypotheses of relationship within Polemoniaceae. The segregation of Ipomopsis and Allophyllum from Gilia was supported by the placement of each in distinct groups separate from a group of four species of Gilia. Several strongly supported groups include genera now placed in different tribes. There was no support for the current separation of temperate Polemoniaceae into two tribes. The tropical genera were resolved as basal and paraphyletic within the family. The family as a whole was monophyletic with no support for the segregate family Cobaeaceae. Sequences of matK, a gene that had not been used previously for phylogenetic analyses, provided a sufficient number of reliable characters for phylogenetic analysis within Polemoniaceae. Pairwise comparisons of matK and rbcL sequences of the same taxa were performed. Sequences of matK varied at an overall rate twice that of rbcL sequences. Substitutions at the third codon position predominated in rbcL sequences, while in matK substitutions were more evenly distributed across codon positions.
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Comparative sequencing of the maturase-encoding chloroplast gene matK has great potential for reconstructing phylogenetic relationships not only within families, but also within genera of land plants. This gene of 1550 bp is easily amplified due to highly conserved, flanking coding regions that include the trnK exons, rps 16, and psbA. Several available sequencing primers also have wide applicability. Parsimony analysis of 45 matK sequences representing Saxifragaceae sensu stricto provides a level of resolution comparable to that obtained via chloroplast DNA restriction site analysis. Furthermore, this analysis suggests relationships among genera and species that are highly concordant with the results of separate analyses of rbcL sequences and chloroplast DNA restriction sites, and with those of combined analyses of these three chloroplast DNA data sets. Parsimony analysis of 31 matK sequences representing all six sections of Gilia (Polemoniaceae) and 10 allied genera provides strong evidence for the polyphyly of Gilia and suggests relationships among sections of Gilia that are highly concordant with a recent ITS sequence analysis of the Polemoniaceae. Our analyses suggest that matK sequences are not strongly biased toward transitions, and the frequency of mutations at the first and second codon positions approach the frequency of mutations in the third codon position.
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identical to print version). TAXON 62 (4) • August 2013: 831 Braun & Heuchert • (2176) Conserve Coniothecium betulinum (2176) Proposal to conserve the name Coniothecium betulinum against Torula stilbospora (Ascomycota). Lectotypus (hic designatus): without locality, on wood, without date, Corda (PRM No. 155699). Torula stilbospora Corda (in Sturm, l.c.) was later confused by Corda (Icon. Fung. 5: 50. 1842) with a fungus nowadays usually treated as "Taeniolella stilbospora (Corda) S. Hughes". Corda's (l.c. 1842: t. 2, fig. 13) second illustration of "Torula stilbospora" is quite distinct from his original drawing (Corda in Sturm, l.c.: t. 46), show-ing a fungus with a more aggregated, granular conidial structure quite distinct from Taeniolella S. Hughes which is in contrast character-ized by forming catenate conidia in firm, oblong-cylindrical chains. This confusion was already realized and discussed by Saccardo (Syll. Fung. 4: 265. 1886). Among original collections deposited in Corda's herbarium (PRM) as Torula stilbospora, there is a single collection without locality and date, but with a small pencil drawing on the envelop agreeing with Corda's original illustration, which represents true type material. This sample has been re-examined and proved to be typical Trimmatostroma betulinum (Corda) S. Hughes (≡ Coniothecium betulinum Corda), i.e., Torula stilbospora agrees with Coniothecium betulinum and is an older heterotypic synonym. Corda (l.c. 1842) cited collections from Neustadtel (locality unclear as at least four towns with this German name are known, two in the Czech Republic [now Dolny Bělá and Jezvé, part of Stružnice], one in Poland [now Nowe Miasteczko] and one in Germany [Saxony]), Senftenberg (Germany) and Prague (Czech Republic). Only the mate-rial from "Neustadtel" collected by Corda (PRM 155698) is preserved and was also re-examined. This material is very sparse, but quite different from the lectotype of Torula stilbo spora. Maintained frag-ments of conidial chains agree well with Corda's (l.c. 1842: t. 2, fig. 13) illustration and represent a genuine Taeniolella, hitherto generally known as T. stilbospora. However, the latter name is misapplied, i.e., the true Taeniolella needs another name. Trimmatostroma betulinum (Corda) S. Hughes is a common, widespread hyphomycete on bark of mostly fallen branches of Alnus, Betula, Corylus, Quercus, Pinus, Populus and Salix species mainly in Europe, but also recorded from Cuba, New Zealand and Turkey, and a widely used name in the mycological literature since its introduction by Hughes did not give any details of the original material on which he based the name C. betulinum, except for the substrate (Betula alba L.), but since only a single collection under this name is deposited in Corda's herbarium, this sample may be considered as holotype material. According to Crous & al. (in Stud. Mycol. 58: 1–32. 2007), this species belongs phylogenetically to the Dermate aceae (Helotiales, Leotiomycetes, Ascomycota). However, the older name Torula stilbospora proved to be a heterotypic synonym of Conio-thecium betulinum, the basionym of Trimmatostroma betulinum, and threatens the latter well-known name. A reallocation of Torula stilbospora, hitherto consistently used in the misapplied combination Taeniolella stilbospora, to Trimmatostroma would undoubtedly cause strong confusion. Hence, a conservation of Coniothecium betulinum over Torula stilbospora is desirable and proposed in order to maintain the current use of Trimmatostroma betulinum.
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The program MODELTEST uses log likelihood scores to establish the model of DNA evolution that best fits the data. AVAILABILITY: The MODELTEST package, including the source code and some documentation is available at http://bioag.byu. edu/zoology/crandall_lab/modeltest.html.
Chapter
Annual or perennial herbs, or subshrubs, rarely shrubs or small trees, usually monoecious, rarely gynodioecious or dioecious. Stems often swollen at the nodes, usually with anomalous secondary growth in older stems, also often occurring in the roots. Leaves opposite, decussate or apparently whorled, very rarely alternate, simple, entire, often connate at the base, sometimes succulent; stipules wanting, more rarely present, then mostly scarious. Flowers commonly protandrous, actinomorphic, very rarely weakly zygomorphic, usually bisexual, rarely unisexual, hypogynous or perigynous, sometimes obdiplostemonous. Inflorescences mostly dichasial, sometimes monochasial, rarely flowers solitary. Sepals (4−)5 or rarely more or fewer, imbricate, rarely valvate, completely free or connate for often most of their length, sometimes subtended by bracts (epicalyx). Petals (4−)5 or rarely more, free, aestivation contorted or rarely imbricate, entire, emarginate, bifid or lacerate, clawed or not, sometimes with coronal scales at the top of the claw, occasionally very small or absent.
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Chromosome numbers of 566 taxa (3852 individuals) of mainly South Swedish vascular plants are published, most of them previously not documented for Sweden. The plant material was collected in the late 50s by Borje Lovkvist and co-workers from natural habitats. The results were not published, although many of Lovkvist's chromosome numbers appear with an asterisk in Weimarck (1963). However, these numbers were largely based on preliminary oral information by Lovkvist and only a little more than half could be verified from Lovkvist's documented material. The entire material (field notebooks, voucher specimens, slides, laboratory journals and drawings) has been critically examined and edited by U-M. Hultgard for the Flora Nordica project. - Flora Nordica Note no. 23.
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Aussi bien par son nombre chromosomique (2n=36) que par certaines de ses particularités morphologiques intéressant la souche, la tige, les feuilles et les graines, le M. valentina (Pau) Mateo & Figuerola offre des affinités avec les espèces de la Section Lanceolatae (Fenzl) Graebner. Un étude de la testa des graines au M.E.B. confirme l'appartenance du M. valentina à cette section (présence de papilles dorsales aplaties- linguiformes). Il ne saurait donc être considéré plus longtemps comme une sous-espèce du M. verna L. Les auteurs proposent de le transférer dans le Section Lanceolatae et de créer pour lui une série nouvelle monospécifique: Series Valentinae Çelebioglu, Favarger & J. M. Monts.
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25 populations from Turkey and one of Syria belonging to theSabulina section of the genusMinuartia have been karyologically examined. New chromosome numbers have been recorded forM. mesogitana andM. hybrida subsp.turcica, and a new variety was found in theM. hybrida complex. The origin of the taxa with n = 23 and n = 35 is discussed.
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Amphitropical distributions of vascular plants in the Western Hemisphere are divided into three groups: bipola or high-latitude, with about 30 species; temperate, with about 130 species; and desert, with a substancial number. Close relationships of this sort can be explained either by divergence from a common tropical ancestor or by origin on one side of the tropics and subsequent migration to the other. The latter is true for the great majority of groups discussed. These may have crossed the tropics in a single jump or by direct migration. The plant communities involved are relatively recent in origin. The amphitropical disjuncts are drawn from relatively few families and mostly are plants that occur in open habitats such as seacoast or seasonally moist places where establishment would be relatively easy. Woody plants and even herbs of closed communities are scarcely represented. Animals by and large do not have analogous amphitropical distributions as they would be expected to if the plants migrated by ...
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A new apetalous species of Minuartia from the high Andes of northwestern Argentina, M. altoandina, is here described and illustrated. At first sight, M. altoandina is morphologically very similar to the European M. sedoides because of the absence of petals and ciliate leaf margin with narrow hyaline teeth; however, from a biogeographical point of view, it will probably be related to the North American M. rossii complex through the morphology of M. austromontana.
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The aim of the present study was to infer a substantially larger, more evenly sampled, phylogenetic tree for Caryophyllaceae in order to more confidently resolve relationships within this clade. This would allow us to evaluate previous classification schemes and to infer the evolution of a number of characters that have figured prominently in higher-level taxonomic treatments. We have inferred a 630-tip phylogeny (ca. 30% of the 2200 species) using maximum likelihood analyses of data from the nuclear ribosomal ITS region and five chloroplast genes and intergenic spacers: matK, ndhF, trnL-trnF, trnQ-rps16, and trnS-trnfM. Our results confirm that subfamily Paronychioideae is paraphyletic at the base of Caryophyllaceae. Alsinoideae and Caryophylloideae together form a clade, within which neither subfamily is monophyletic. With only a few exceptions, our results support the tribal classification presented by Harbaugh & al. (2010). In agreement with other recent studies, it appears that many of the larger genera are not strictly monophyletic. Our results imply that the first Caryophyllaceae had stipules, free sepals, small apetalous flowers with few stamens, and single-seeded indehiscent or irregularly dehiscing utricles. Stipules were lost along the branch to the Alsinoideae-Caryophylloideae clade, and the evolution of a tubular calyx marks Caryophylloideae. The evolution of petals, 10 stamens, and capsule fruits is inferred to have taken place along the branch subtending a clade that includes Sperguleae (mostly containing former members of Paronychioideae) and the remainder of Caryophyllaceae. As this previously unnamed major group is both well-supported in molecular phylogenetic studies and marked by clear-cut apomorphies, we propose the name Plurcaryophyllaceae for this clade and provide a phylogenetic definition.
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Minuartia dirphya from Mt Dirphys (Evvia) is described as a species new to science. It is closely related to M. wettsteinii from Mt Thriptis (East Kriti) and M. parnonia from Mt Pamon (South Peloponnisos). The three narrow endemic species form a well-defined group, the Minuartia wettsteinii-group, within Minuartia ser. Laricifoliae Mc Neill. Additions to the descriptions, chorological and ecological data for M. wettsteinii and M. parnonia are provided. The morphological characters of the group and the differences to the other species of ser. Laricifoliae are discussed. A key to the species, illustrations of the taxa and their seeds, chromosome numbers and data on their geographical distribution are also provided.
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The importance and abundance of cryptic species among invertebrate taxa is well documented. Nowadays, taxonomic, phylogenetic and conservation biological studies frequently use molecular markers to delineate cryptic taxa. Such studies, however, often face the problem of the differential resolution of the molecular markers and techniques involved. This issue is explored in the present study of cryptic taxa within the terrestrial slug complex Arion subfuscus/fuscus in continental north-west Europe. To this end, morphological, allozyme and mitochondrial 16S rDNA sequence data have been jointly evaluated. Using allozyme data and gonad type, two distinct groups were consistently delineated, even under sympatric conditions. The 16S rDNA data strongly supported both those groups and even suggested the presence of three distinct taxa within one of them. However, in view of: (1) the allopatric distribution of three OTUs, (2) the lack of allozyme or morphological differentiation, and (3) the extremely high degree of intraspecific mtDNA variation reported in pulmonate gastropods, they are, for the time being, not regarded as valid species under the biological species concept. By means of 16S rDNA and allozyme data, the position of type and topotype material of A. subfuscus s.s. and A. fuscus relative to the newly defined OTUs was determined, thus clarifying the nomenclature of this species complex. Additionally, gonad type proved to be a useful character for distinguishing the two species in north-west Europe. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 23–38.
Article
Caryophyllaceae is a principally holarctic family including around 2200 species often classified into the three subfamilies Alsinoideae, Caryophylloideae, and Paronychioideae. Complex and possibly homoplasious morphological characters within the family make taxa difficult to delimit and diagnose. To explore part of the morphological evolution within the family, we investigated the phylogeny of the Caryophyllaceae by means of analyzing plastid and nuclear sequence data with parsimony and Bayesian methods. We describe a mode of tracing a stable phylogenetic signal in ITS sequences, and a significant common signal is shared with the plastid data. Parsimony and Bayesian analyses yield some differences in tree resolution. None of the subfamilies appear monophyletic, but the monophyly of the Caryophylloideae is not contradicted. Alsinoideae are paraphyletic, with Arenaria subg. Eremogone and Minuartia subg. Spergella more closely related to the Caryophylloideae. There is strong support for the inclusion of Spergula-Spergularia in an Alsinoideae-Caryophylloideae clade. Putative synapomorphies for these groupings are twice as many stamens as number of sepals and a caryophyllad-type of embryogeny. Paronychioideae form a basal grade, where tribe Corrigioleae are sister to the rest of the family. Free styles and capsules with simple teeth are possibly plesiomorphic for the family.
Moehringia 13. Seeds exarillate 14 14. Flowers strongly perigynous
  • ........................................ Seeds Arillate.................................................................................. S And C Asia ).................................... Styles
Styles (carpels) 3–5.......................................... 22 13. Seeds arillate..................................... Moehringia 13. Seeds exarillate............................................... 14 14. Flowers strongly perigynous (S and C Asia)............................................................... Thylacospermum 14. Flowers weakly perigynous or hypogynous............. 15
19 16. Stamens 2 Arenaria 17. Capsule 2-toothed Bufonia 18. Plants minute, moss-like; flowers solitary (Chile) Reicheëlla Pax 18. Plants rush-like herbs or subshrubs; flowers not solitary (Mediterranean and Irano-Turanian regions)
  • ...................................................................................................................................................................................................................................................................................... Sepals
Sepals and petals 5........................................... 19 16. Stamens 2..................................................... 17 16. Stamens 3–8.................................................. 18 17. Capsule 4-valved or -toothed...................... Arenaria 17. Capsule 2-toothed.................................... Bufonia 18. Plants minute, moss-like; flowers solitary (Chile)........................................................... Reicheëlla Pax 18. Plants rush-like herbs or subshrubs; flowers not solitary (Mediterranean and Irano-Turanian regions).... Bufonia 19. Capsule with 2 valves or teeth................ Lepyrodiclis 19. Capsule with 4 valves or teeth............................. 20 20. Ovules and seeds numerous....................... Arenaria 20. Ovules 2(–5);
Reli-quiae haenkeanae; seu Descriptiones et icones plantarum quas in America meridionali et boreali Contributions to a chromosome atlas of the New Zealand flora – 25 miscellaneous species
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Contribution à la cytotaxono-mie du genre Minuartia L. (Caryophyllacées) en Turquie et dans quelques régions voisines
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